First published online August 23, 2004
Journal of Experimental Biology 207, 3289-3298 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01155
Grouping of visual objects by honeybees
Shaowu Zhang1,*,
Mandyam V. Srinivasan1,
Hong Zhu1 and
Jason Wong2
1 Centre for Visual Sciences, Research School of Biological Sciences,
Australian National University, Canberra, ACT 2601, Australia
2 School of Molecular and Microbial Sciences, University of Sydney, Sydney,
Australia

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Fig. 1. Illustration of experiments showing (A,B) the multiple-choice maze
apparatus and (C) the four groups of stimuli that were used. See text for
details.
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Fig. 5. Results of four control tests to check for the possible use of olfactory
cues. Bees trained in the category discrimination experiments of
Fig. 2 were presented with
tests in which the sample as well as the four test stimuli were identical (in
this case, they were grey-level versions of F1). The feeder was placed behind
one of the test stimuli, as shown by the circular symbol. The bees chose
randomly among the four stimuli, indicating that they were not using olfactory
cues to make their discriminations in our experiments.
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Fig. 2. Results of (A) learning tests and (B,C) Type 1 transfer tests. For each
group, the bars show the relative preferences for the four test stimuli when
the sample was a star-shaped flower, a circular flower, a plant stem or a
landscape, as shown underneath the abscissa. In each panel, N denotes
the number of bees that were tested in each experiment. Asterisks denote
statistically significant differences from the random choice level of 25%
(broken horizontal lines). ***P<0.001;
**P<0.01; *P<0.05. Black
asterisks refer to levels that are significantly greater than 25%, and red
asterisks to levels significantly lower than 25%. The circles denote no
significant difference from 25%. Values are means ±
S.E.M. of the data. In each case, the bees
are able to learn to choose the test stimulus that matches the sample.
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Fig. 3. Results of Type 2 transfer tests examining the ability of bees, having
encountered a sample stimulus from Group 1, to choose a test stimulus of the
same category in (A) Group 2, (B) Group 3 and (C) a novel Group 4. In each
case, the bees are able to learn to choose the test stimulus that belonged to
the same category as the sample. Details as in
Fig. 2.
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Fig. 4. Transfer tests examining the ability of bees, having encountered a sample
stimulus from Group 3, to choose a test stimulus of the same category in Group
3*, which is composed of grey-level versions of the stimuli in
Group 3. Bees are clearly able to perform this task. Details as in
Fig. 2.
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Fig. 6. Power spectra of patterns in each of the four groups of stimuli used in the
experiments. Spatial frequency ranges from -2.22 cycles cm-1 to
+2.22 cycles cm-1 along the u and v axes, where
the unit spatial frequency represents a period equal to the diameter of the
full image.
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© The Company of Biologists Ltd 2004